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Should tree invasions be used in treeless ecosystems to mitigate climate

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Article  in  Frontiers in Ecology and the Environment · May 2021

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 REVIE WS    1

Should tree invasions be used in treeless

ecosystems to mitigate climate change?
Martin A Nuñez1,2*, Kimberley T Davis3, Romina D Dimarco4, Duane A Peltzer5, Juan Paritsis6, Bruce D Maxwell7, and
Aníbal Pauchard8,9

Intentionally allowing or promoting invasion by non-native trees into areas characterized by treeless vegetation could contribute
to climate-­change mitigation by increasing carbon (C) sequestration. In some areas of the world, incentives exist to retain invasive
non-native trees in natural systems as a mechanism for increasing ecosystem C storage and reducing atmospheric carbon dioxide
levels. Although this novel opportunity for C sequestration holds appeal, such an approach is problematic for several reasons:
(1) invasive trees do not always increase net C sequestration due to greater occurrence of fire or reduced soil C; (2) lower albedo in
invaded areas can increase absorption of solar radiation, thereby offsetting potential C sequestration; and (3) tree invasions often
also have negative effects on biodiversity, economic opportunities, and water yield. Such drawbacks are sufficient to raise doubts
about the widespread use of non-native tree invasions in treeless areas as a tool to ameliorate climate change.

Front Ecol Environ 2021; doi:10.1002/fee.2346

A ction is needed to limit the magnitude and rate of climate

   change, one of the most crucial problems facing humankind
(IPCC 2014). Despite their potential negative impacts on biodi-
versity and human welfare, non-native invasive woody plants have
been proposed as a tool for climate‐­change mitigation through
In a nutshell:
• Non-native tree invasions can increase carbon (C) se-
questration in some ecosystems; consequently, unmanaged
incursions of introduced tree species are often considered
a potential opportunity for climate‐­change mitigation
• The overall effect of tree invasions on climate‐­ change
mitigation is poorly understood, but evidence suggests
net negative effects in some instances
• Tree invasions can alter fire regimes, soil C sequestration,
and light absorption, all of which influence the impact
invaders may have on climate regulation
• Overall, the detrimental impacts of tree invasions on bio­
diversity, economic opportunities, and water yield may
offset any positive effects on C sequestration
• Managers or organizations considering using non-native
trees for C sequestration should take into account the
diverse problems associated with plant invasions
1
Grupo de Ecología de Invasiones, INIBIOMA, CONICET, Universidad
Nacional del Comahue, San Carlos de Bariloche, Argentina; 2Department
of Biology and Biochemistry, University of Houston, Houston, TX
*
(nunezm@gmail.com); 3Department of Ecosystem and Conservation
Sciences, University of Montana, Missoula, MT; 4Grupo de Ecología de
Poblaciones de Insectos, IFAB (INTA-­CONICET), San Carlos de
Bariloche, Argentina; 5Manaaki Whenua Landcare Research, Lincoln,
New Zealand; 6Laboratorio Ecotono, INIBIOMA, CONICET,
Universidad Nacional del Comahue, San Carlos de Bariloche, Argentina;
7
Land Resources and Environmental Sciences Department, Montana
State University, Bozeman, MT; (continued on last page)
enhanced carbon (C) sequestration (Liao et al. 2008; Pejchar and
Mooney 2009). For example, in some countries (eg New Zealand),
monetary incentives in the form of C credits are used to discour-
age the removal of non-native invasive trees (Mason et al. 2017).
Interest in this approach is growing because C sequestration by
trees is a fundamental tool to mitigate climate change (Bastin et al.
2019; Griscom et al. 2017) and because tree invasions are increas-
ingly common (Richardson et al. 2014). In many situations, inva-
sive trees are not at present being retained to mitigate climate
change, but employing them as C sinks in the future could under-
mine broader efforts to control invasives. Allowing tree invasions
for the purpose of C sequestration may have limitations and unin-
tended consequences similar to intentional tree planting for C
storage (eg Holl and Brancalion 2020), but may also come with
unique challenges. Leaving tree invasions intentionally uncon-
trolled as C sinks could be problematic for at least two primary
reasons: tree invasions may not sequester more C long‐­term com-
pared to the ecosystem they replace, and tree invasions may trig-
ger severe economic and environmental impacts.
Invasive trees often grow faster in areas in which they have
been introduced than in their native ranges (Parker et al. 2013;
Davis et al. 2019). Moreover, trees invading treeless ecosystems
potentially transform these areas from low aboveground C
sequestration to areas of high aboveground C accumulation.
Although this factor increases the appeal of using tree inva-
sions as C sinks, maintaining tree invasions as C sinks in tree-
less ecosystems may generate negative impacts that offset or
even outweigh the potential benefits.
Understanding how climate can be affected by tree
invasions
In forested ecosystems, C sequestration potential by invasive
woody species may be low due to the presence of native

© The Ecological Society of America Front Ecol Environ doi:10.1002/fee.2346

 2   RE V IE WS
Figure 1. Diagram of the problems and benefits of woody invasions described in the main pruning lower branches), which can limit
text, including changes in fire regimes, aboveground and belowground carbon (C) sequestra- vertical fuel connectivity; however, tree inva-
tion, water use, species diversity, and albedo.
trees. Tree invasions into treeless ecosystems (eg grasslands,
shrublands) are widespread (Rundel et al. 2014) but may
not enhance C sequestration because of increased fire risk
and reduced soil C. In addition, because the overall aim
of C credits is to minimize global warming, woody species
invasions may result in lower albedo, raising land surface
temperatures. Here, we present and discuss the evidence
Panel 1. A worked example of pine invasion, carbon, and fire from the Southern Hemisphere
Native to North America, lodgepole pine (Pinus contorta) has been
introduced into several regions of the Southern Hemisphere, where
it quickly invades open ecosystems and grows faster than in its
native range (Taylor et al. 2016a), therefore allowing for rapid rates
of aboveground carbon (C) sequestration. However, lodgepole pine
invasions in Argentina, Chile, and New Zealand were found to greatly
increase fuel loads, especially in treeless areas, which is expected to
cause more intense fires than in uninvaded stands (Taylor et al. 2017;
Paritsis et al. 2018). To quantify how much C would be released into
the atmosphere if invaded and uninvaded grassland and shrubland
Front Ecol Environ doi:10.1002/fee.2346
MA Nuñez et al.
suggesting that invasion of non-native tree
species into treeless ecosystems is a less than
ideal mechanism for C sequestration, and that
it can have detrimental, unintended environ-
mental impacts, even including promotion of
positive climate‐­change feedbacks (Figure 1).
Changes in fire regimes
Aboveground biomass accumulation due to
tree invasion increases fuel loads and alters
fuel distribution, which in turn modifies fire
regimes and enhances fire risk (eg Mandle
et al. 2011; Souza‐­Alonso et al. 2017; Castro‐­
Díez et al. 2019). For example, dense invasions
of broad‐­leaved paperbark (Melaleuca quin-
quenervia) into Florida prairies and wetlands
have caused a shift in fire regime from low
to high intensity (Mandle et al. 2011). In
fuel‐­limited systems like the Patagonian
steppe of South America, woody invasions
increase both fuel loading and connectivity,
leading to increased fire intensity and severity
(Taylor et al. 2017; Paritsis et al. 2018). In
contrast, some invasive trees have traits that
reduce fire spread in areas with frequent‐­fire
regimes (eg Stevens and Beckage 2009).
Plantations of non-native trees are typically
managed to improve wood quality (eg by
sions into grasslands or shrublands have lower
crown base heights that connect surface veg-
etation to the canopy, thereby increasing the risk of crown
fire (Paritsis et al. 2018). Fire season length and fire activity
are widely projected to increase in many parts of the world
due to climate warming (Jolly et al. 2015), making stands
of invasive trees (particularly flammable species, such as
Pinus and Eucalyptus) more likely to burn and release stored
C back into the atmosphere (Panel 1 and Figure 2).
sites in New Zealand and Patagonia were burned, we used fuel
loads collected across a lodgepole pine invasion gradient (Taylor et
al. 2017) and the First Order Fire Effects Model (Reinhardt 2003) to
simulate C emissions from a wildfire. We found significantly higher
simulated C emissions in invaded as compared to uninvaded plots at
three of four study sites (two in New Zealand, one each in Argentina
and Chile; ANOVA and post-­hoc Tukey’s tests). Stands of invasive
pines that were ~10 years old were found to release much larger
amounts of carbon dioxide into the atmosphere under fire simulations
than uninvaded plots (Figure 2).
© The Ecological Society of America
 Non-native tree invasions and climate change REV IE WS    3

Reductions in soil C (a)

Although increased aboveground C storage following woody
species invasion is well documented, much less is known
about belowground storage. This is surprising because soil
C storage is the primary component of global C sequestra-
tion, with two‐­to threefold more C stored in soils than in
terrestrial vegetation (Houghton 2007). A meta‐­analysis across
invasive species and ecosystems revealed that soil C can
increase slightly following invasion (Liao et al. 2008), but
recent experimental evidence suggests that non-native species
reduce soil C through interactions with herbivores and soil
biota (Waller et al. 2020). Other studies focused on woody
species suggest that soil C can either increase or decline
after invasions (Jackson et al. 2002) due to changes in root-
ing depth, associated soil biota, or lower C inputs from
resident species. For instance, co‐­invasion of ectomycorrhizal
tree species, which are globally important invaders (eg all (b)

Pinaceae and Eucalyptus, and species in the Salicaceae and

Acacia), and their symbionts can increase rates of nutrient
cycling and oxidation of C pools compared to native myc-
orrhizal forms, thereby reducing soil C in comparison to
native vegetation (Farley et al. 2004; Dickie et al. 2014).
C accumulation in soils is driven by numerous and complex
processes that are both directly and indirectly influenced by
invasive trees (Sapsford et al. 2020) through, for example, differ-
ences in biomass allocation or accretion of the invader itself,
longevity of the non-native species, or alteration of litter quality
and quantity to the soil subsystem and therefore C and nutrient
cycling (eg Castro‐­ Díez et al. 2014). The relatively rapid
increase in aboveground biomass of invaders (Liao et al. 2008)
commonly exerts important indirect effects on ecosystem C by
altering the composition, diversity, and function of resident
vegetation (Wardle and Peltzer 2017; Davis et al. 2019). Overall,
the assumption that tree invasions will promote soil C levels Figure 2. Simulated carbon dioxide emissions as a function of time since
may not be true in all cases, underscoring the need for more lodgepole pine (Pinus contorta) invasion (years) and in comparison to unin-
comprehensive species‐­and system‐­specific information. vaded plots in grassland systems, at sites in Chile (a) and New Zealand (b).
Horizontal lines within boxes depict median values, boxes represent the
interquartile range (25th–­75th percentiles), whiskers (vertical lines) repre-
Reduction in albedo
sent 1.5×interquartile range, and solid circles depict outliers. Asterisks
Increased forest cover in the temperate and cold regions of highlight groups that were significantly higher (P < 0.05) than the unin-
the world can produce a net warming of the atmosphere even vaded plots based on post-­hoc Tukey’s tests.
under scenarios of C accumulation due to altered surface
albedo (Arora and Montenegro 2011; Davies‐­ Barnard et al. managed stands). Other studies have reported even greater
2014; Kreidenweis et al. 2016). Changes in albedo are funda- differences; for instance, evergreen plantations and grasslands
mental to understanding the net effect of tree invasions on were characterized by albedos of 0.12 and 0.19, respectively
global warming, especially in treeless areas. Because of the (that is, 88% and 81% of the sunlight received by these land‐­
vast extents of the planet’s terrestrial surface that could poten- cover types would be absorbed) (Schaeffer et al. 2006). It was
tially be occupied by non-native tree species, changes in reflec- beyond the scope of this article to calculate temperature increases
tion could undermine the overall goal of using invasive woody that could result from such invasions, but these findings illus-
species as a tool to mitigate climate change. To illustrate, we trate that changes in albedo of this magnitude could contribute
observed a 20% reduction in albedo in a native steppe in to shifts in global temperature if they occur over large spatial
Chile within ~10 years of pine invasion (Figure 3); if unman- scales, possibly producing a net rise in temperature even under
aged, it is likely that the invaded area will eventually attain a scenario of high C accumulation (Davies‐­ Barnard et al.
albedo levels similar to those of nearby plantations (or possibly 2014; Kreidenweis et al. 2016). Although albedo can be affected
even lower, given the higher tree density in invaded versus by complex factors (eg cloud cover), a change in albedo of

© The Ecological Society of America Front Ecol Environ doi:10.1002/fee.2346

 4   RE V IE WS
Figure 3. Total shortwave albedo (mean ± standard deviation) obtained
from Landsat 7 products for the Coyhaique area, Chile, for a Patagonian
steppe, a dense but recent (~10-­year-­old) lodgepole pine (P contorta)
invasion in the steppe, and a lodgepole pine plantation in an adjacent area.
Albedo was calculated based on 20 points for each vegetation type using
equations described in Liang (2001).
0.01 at a global scale would have a warming effect equal to
that generated by a doubling of the current amount of carbon
dioxide in the atmosphere (Wielicki et al. 2005).
Impacts of tree species invasion beyond
climate‐­change mitigation
Tree species invasions into treeless areas may have multiple
ecosystem‐­scale effects, many of which –­such as altering C
Panel 2. A practical example of the potential conflicts and concerns of using tree invasions for C sequestration in New Zealand
Species of Pinaceae underpin the plantation forestry industry in New
Zealand but are also widely naturalized biological invaders. For example,
wood exports from pines comprise ~99% of total log volume exports
and are the fourth largest industry nationally (NZMPI 2019). On the
other hand, at least 14 species of Pinaceae are considered to consti-
tute a serious weed problem on ~1.8 million ha (Froude 2011; Hulme
2020). As a consequence, government agencies, land managers, and
communities currently spend in excess of NZ$15 million annually on
management. Overall, there are nontrivial costs and benefits of non-
native pines in New Zealand that have generated much debate over
their management, both as a resource and as invaders. One of these
issues is whether invasive pines should be retained in some areas for
C sequestration. There is active debate on the pros and cons of using
invasive trees for C sequestration. This debate spills over directly into C
sequestration policy through New Zealand’s Emissions Trading Scheme
(ETS; www.mfe.govt.nz/ets), which is intended to promote more envi-
ronmentally sustainable management by C emitters paying for C credits
from entities that remove greenhouse gases. At present, New Zealand
is the only country so far to include plantation forests as full partici-
pants in an ETS (Evison 2017). A large number of plantations that were
Front Ecol Environ doi:10.1002/fee.2346
MA Nuñez et al.
sequestration –­can be pervasive (Panel 2 and Figure 4).
First, these invasions often lower the abundance and diversity
of native species (Pyšek et al. 2012; Davis et al. 2019) and
induce shifts in soil biotic communities and nutrient cycling
(Le Maitre et al. 2011; Castro‐­Díez et al. 2019), resulting in
fundamental and persistent changes to ecosystems. Second,
greater aboveground C sequestration is associated with
increased water use and consequently lower water yield in
catchments. Global studies show afforestation of grasslands
or shrublands reduces streamflow and runoff by 40–­ 75%
(Farley et al. 2005; Jackson et al. 2005), which can restrict
water availability to urban areas (Pejchar and Mooney 2009)
and exacerbate problems of surface and groundwater avail-
ability in dry regions (Le Maitre et al. 1996, 2000). In addi-
tion, decreased soil organic matter beneath invasive pines
can reduce soil water retention relative to native grasslands
(Farley et al. 2004). Greater water use can also interact with
other factors, for example, by increasing fire risk. Third,
woody species invasions can have severe negative social and
economic impacts across diverse sectors, such as plantation
forestry, tourism, and sheep and cattle ranching (Ledgard
2001; Le Maitre et al. 2011; van Wilgen and Richardson
2012; Castro‐­Díez et al. 2019). Heavily invaded areas cannot
be used for other activities without the removal of trees,
which is often unaffordable for landowners (Nuñez et al.
2017). For example, at sites in New Zealand, the cost of
removing invasive pines ranges from NZ$1–­ 50 per hectare
for sparse invasions to more than NZ$2500 per hectare for
dense invasions, and management is often repeated within a
site before a different land use is feasible. Other major eco-
nomic impacts include the reduction of surface streamflow,
established in the 1980s through afforestation incentives are now due
for harvest, and this has created an urgent need to replace forests that
can rapidly sequester C to meet international climate obligations, such
as the 2015 Paris Agreement. Post-­1989 forests registered in the ETS
can be liable for deforestation by removing invasive trees; likewise, pre-­
1990 forests require a “tree weed” exemption if subject to the ETS. The
1989/1990 cutoff in managing forests for C sequestration means that,
in some instances, there are ~30 years of potential invasion that either
can be partially claimed under ETS or require exemption to allow for
weed management. Although tree invasions can be considered under
some circumstances to provide benefits for C sequestration, the trade-­
offs involved with ongoing invasion and negative impacts mean that
retaining invasive trees is not generally considered acceptable practice
over the long term (Edwards et al. 2020). However, invasive trees are
retained in some long-­invaded or remote areas where management is
considered intractable or unaffordable, but these are not included in
the ETS. Ultimately, this example highlights that the interplay between
management of biological invaders, policy, and practice underpins deci-
sions for when and where tree invaders are removed or retained (Hulme
2020).
© The Ecological Society of America
 Non-native tree invasions and climate change
for instance after the introduction of acacias in South Africa,
which resulted in losses equivalent to over US$200 million
and exacerbated social conflicts in the region (De Wit et al.
2001; Shackleton et al. 2018). Tree invasions can also alter
landscapes and their aesthetics, driving shifts in intrinsic,
tourism‐­ based, and recreational values (Castro‐­ Díez et al.
2019). Collectively, these impacts and the shifts they produce
(eg in fire regimes or water cycles) suggest that tree invasions
can cause major environmental and social problems (Kull
et al. 2018). How the total economic costs or benefits of
biological invaders can be quantified adequately across their
multiple effects on environmental, social, and economic factors
remains unresolved (Bartkowski et al. 2015).
When would it be justified to use tree invasions as a
climate‐­change mitigation effort?
Although leaving woody species invasions unmanaged to act
as C sinks has several disadvantages, this approach may be
a viable option for species that have a net positive effect on
C sequestration over the long term but only minor adverse
impacts on biodiversity and ecosystem services. Some non-
native woody species are more invasive than others, which
can be explained by their characteristics. For example, lodge-
pole pine (Pinus contorta) is highly invasive in the Southern
Hemisphere due in part to its relatively small seed size and
early age of reproduction (Richardson and Rejmánek 2004),
making removal of this species essential regardless of its value
to C sequestration because it can quickly spread into areas
where it is unwanted; in addition, large‐­scale removal is costly
and can produce negative soil legacy effects (Nuñez et al.
2017; Dickie et al. 2014). Similarly, invasions by some Acacia
spp have negative ecological or economic impacts (Souza‐­
Alonso et al. 2017) that far exceed the benefits of their C
sequestration, such as reductions in water availability for
crops and urban areas (De Wit et al. 2001). In contrast,
other types of conifers (eg cypress [Cupressus spp]) do not
readily spread in some areas (Richardson and Rejmánek 2004),
and therefore may be preferable to more invasive species.
Retaining invasive tree species may also be considered if
populations can be contained or managed within specific
sites. Some woody species only become invasive in specific
environments (eg degraded pastures) and may be passive
bystanders as opposed to active drivers of ecological change
(MacDougall and Turkington 2005); spread of these species
may be more easily controlled, making them better suited
for C sequestration purposes. At present, however, few man-
agement options exist for the efficient removal of invasive
woody species, and containment requires ongoing manage-
ment of buffers and surveillance to confirm effectiveness
(Panetta 2012). Furthermore, removal of woody non-natives
will not necessarily restore the ecosystem to its previous
state (Sapsford et al. 2020) but instead may induce shifts
toward different community compositions and ecosystem
© The Ecological Society of America
REV IE WS    5
Figure 4. Pine invasions create landscape-­level changes in both fire
regimes and ecosystem properties, including C sequestration. This image
shows lodgepole pine (P contorta) and Austrian pine (Pinus nigra) invading
native tussock grasslands in the Southern Alps of New Zealand. Any C
sequestration driven by pine invasion may be short-­lived due to increased
fire disturbance or large-­scale management to remove these invasive
trees.
processes, and in some cases additional invasions by other
non-native species (Nuñez et al. 2017). Overall, the manage-
ment costs of retaining invasive tree species can be consider-
able, and these costs must be balanced against the benefits
for climate mitigation or other services. Yet invasions should
still be used with caution and continuously monitored given
the ongoing and long‐­term potential for non-native species
to both invade and, in some cases, fundamentally alter eco-
systems (eg Strayer et al. 2006; Hulme 2020).
Are there more sustainable alternatives to using
invasions?
Native species that spread into new ecosystems can also
be used for C sequestration (Simberloff et al. 2011).
Expansion of native tree species is a widespread phenom-
enon and may have fewer negative consequences for the
environment than non-native invasive species given their
coevolutionary history with the invaded community.
However, invasion of native species into adjacent areas
(that is, woody encroachment) can also result in loss of
biodiversity (Jackson et al. 2002; Taylor et al. 2016b),
shifts in fire regime (Ratajczak et al. 2014), and possibly
lower C sequestration. Evaluation of leaving native invaders
uncontrolled to serve as C sinks must therefore also take
ecosystem impacts into account.
Native and non-native tree species in planted forests with
clear commercial value and defined management plans may
Front Ecol Environ doi:10.1002/fee.2346
 6   RE V IE WS
be a viable option for use as C sinks. Commercial value may
derive from firewood, timber, or a non‐­wood forest product
used in the food, chemical, or pharmaceutical industries
(Rodrigues‐­Corrêa et al. 2012; Hulme 2020). In these cases,
it is important to assess how invasive the commercial tree
species is, given that it could escape into areas where it can-
not be harvested commercially (Nuñez et al. 2012).
Management plans of plantations should thus consider the
control of all individuals that spread beyond the original
plantation stands (eg through inclusion into Forest
Stewardship Council standards).
Restoration of areas previously occupied by trees has been
promoted as a key tool to increase C sequestration at a global
scale (Griscom et al. 2017; Bastin et al. 2019). This alternative
is ideal if the species included in the restoration were present
historically (ie native). Large amounts of C can be stored
through forest restoration, and this should be a priority given
the combined benefit of C sequestration and the restoration of
natural ecosystems.
How can management decisions be improved?
Economic and technical resources for controlling invasive
species are often limited. In these instances, a detailed anal-
ysis of C sequestration and other impacts –­taking into
account soil C, fire activity, albedo, biodiversity, and water
use –­should be considered before the decision whether to
retain an invasive species is made. Although invasions may
provide net C accumulation, there are still no effective man-
agement options available, as with pine invasions in different
parts of the world (Nuñez et al. 2017). In these scenarios,
available methods for evaluating impacts (eg economic impact
classification of alien taxa [EICAT], socioeconomic impact
classification of alien taxa [SEICAT]; Bacher et al. 2018)
should be used to produce objective impact assessments,
which can then be used to determine the costs and benefits
of different management options.
In cases where effective management options are availa-
ble, net C sequestration and positive impacts on mitigating
climate warming should be determined; these positive
effects of C sequestration must be considered along with
negative effects, such as impacts on local economies or bio-
diversity, when deciding whether the affected area should be
prioritized for management. For example, funds available for
climate‐­change mitigation can be allocated toward removal
of invasive species that promote climate warming due to a
reduction in long‐­term C sequestration or in albedo. As an
example, several areas in New Zealand that are managed for
pine invasions under a national wilding conifer control pro-
gram are now being considered for subsequent afforestation
under a different national initiative, the One Billion Trees
Programme (Te Uru Rākau 2018), which uses both native
trees and non-native tree species with low risk of invasion.
Such an approach both reduces the current and future risks
Front Ecol Environ doi:10.1002/fee.2346
MA Nuñez et al.
of tree invasion and supports a shift toward afforestation for
climate mitigation.
When contemplating whether to remove or retain invasive
trees it will be crucial to apply objective decision‐­making tools.
The decisions might be similar to those made when consider-
ing managed relocation (or assisted migration), for which
multicriteria decision frameworks have been developed (eg
Richardson et al. 2009). In this regard, there are important sci-
entific, regulatory, and ethical challenges that must be taken
into account (see Schwartz et al. 2012). Experimental evidence
concerning impacts and C accumulation is frequently lacking,
yet such information is fundamental for decision making and
adaptive management under global change scenarios.
Conclusions
Determining whether an invasive species can and should be
retained as a C sink to help mitigate climate change involves
consideration of aspects beyond its aboveground C storage
capacity. Many factors play a role in decision making con-
cerning the use of invasive woody species as climate‐­change
ameliorators. Climate change and biological invasions are com-
plex problems requiring solutions that incorporate scientific,
economic, and social considerations. The objective of this review
was to show that woody species invasions are rarely effective
or desirable in mitigating climate change because their effects
on C sequestration are not always positive, and they can have
a range of detrimental impacts on ecosystems.
Acknowledgements
We thank P Kardol and C Iversen for helpful comments
on early versions of the manuscript. MAN was supported
by PICT 2016‐­1412 and PICT 2018 329; DAP was supported
by the Winning Against Wildings research program funded
by the New Zealand Ministry of Business Innovation and
Employment; AP was funded by CONICYT PIA AFB170008;
and MAN and AP were funded by NERC NE/S011641/1.
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8
Laboratorio de Invasiones Biológicas, Facultad de Ciencias Forestales,
Universidad de Concepción, Concepción, Chile; 9Institute of Ecology and
Biodiversity, Santiago, Chile
© The Ecological Society of America